Review of Chinese Spiraea (Rosaceae, Spiraeoideae) withSimple Inflorescences

Roman Businsky

Silva Tarouca Research Institute for Landscape and Ornamental Gardening (RILOG),252 43 Pruhonice, Czech Republic. businsky@vukoz.cz

ABSTRACT. Several morphotypes of simple inflores- with simple inflorescences from Yunnan and Taiwancences occurring in the genus Spiraea L. (Rosaceae), that had been hitherto confused are newly distin-widely used for classification and determination, but guished here. One species from each pair is foundnot uniformly interpreted in the literature, are either as a new species or as one neglected for thecorrectly defined. Taxonomically, S. compsophylla Chinese flora. Inflorescence type has traditionallyHand.-Mazz. is synonymized with the imperfectly been the most important character used to classify theknown S. calcicola W. W. Sm., whose amplified genus Spiraea, but a certain confusion in itsdescription and illustration are added. A newly interpretation exists. Two principal inflorescencedistinguished species, hitherto confused with the types are found in this genus, distinguished as thoseformer name, is described from Yunnan Province in with 1-flowered pedicels (simple inflorescences) orChina and illustrated as S. adiantoides Businsky, those with branched pedicels (compound inflores-with its assignment to section Glomeratae Nakai. Two cences). Both types are of indeterminate (racemose)imperfectly known and confused species of Spiraea character, i.e., the lower flowers (for compoundsect. Chamaedryon Ser. from Taiwan, S. tarokoensis inflorescences, those of each segment) open first.Hayata and S. tatakaensis I. S. Chen, are carefully Although these two inflorescence types discriminateddescribed, illustrated, and compared with six similar infrageneric division within the genus (for the historyChinese species. A new variety, S. lasiocarpa var. of its classification, see Businsky & Businska, 2002),villosa Businsky, is described from western Sichuan recent molecular analyses (Potter et al., 2007) revealProvince, accepting the concept that S. lasiocarpa that neither inflorescence type is monophyletic.Kar. & Kir., described from today’s eastern Kazakh- Maximowicz (1879: 173), in his comprehensivestan and hitherto neglected in Chinese floras, is the classification of the genus, concisely defined inflo-correct name for the Chinese taxon S. mongolica rescences of true spiraeas as ‘‘flores haud vereMaxim. Two imperfectly known taxa from southwest- racemosi, pedicellis inaequilongis.’’ He indicated theern Sichuan, S. muliensis T. T. Yu & L. T. Lu and S. simple type as ‘‘corymbi vel fasciculi pedicellis 1-daochengensis L. T. Lu, were found to be conspecific floris’’ and the compound type as ‘‘corymbi velafter comparison of their respective holotypes and a paniculae compositae.’’ Schneider (1905: 449, 450,newly found population; the former is accepted as the 454) described inflorescences of the subgenuscorrect name. Spiraea sericea Turcz. is accepted only Euspiraea C. K. Schneid. (the genus Spiraea byat the rank of the variety as S. media Schmidt var. today’s concept) in German as ‘‘6 scheindoldig,sericea (Turcz.) Maxim. A key to the Chinese species einfach doldentraubig oder 6 zymos verastelt, nieof Spiraea with simple inflorescences is newly deutlich gestreckte Trauben.’’ He divided the simplecompiled, encompassing 40 species. type into one group with ‘‘Blutenstand doldig,Key words: China, IUCN Red List, Rosaceae, sitzend’’ and another group with ‘‘Blutenstand stets

Spiraea. 6 doldentraubig, gestielt.’’ Rehder (1927: 333,1940: 328) indicated the simple types similarly as

Seventy species of the genus Spiraea L. (Rosaceae, ‘‘sessile umbels’’ and ‘‘umbel-like racemes on leafySpiraeoideae) have recently been reported from branchlets.’’ Of these three cited authors, Maximo-China, including Taiwan, in the Flora of China wicz’s definition is the most accurate. The simpletreatment (Lu & Crinan, 2003). However, certain inflorescence type found in Spiraea is alwaysoversights exist in the treatment of this taxonomically determined by its lateral position on the previouscritical genus. These concern certain characters in year’s shoots. They differ only in the length of thesome imperfectly known species, as well as the inflorescence axis, the continuing peduncle, andwording used in certain group leads in the key to short branchlet. This length difference can bespecies. As a result of a field study, two species pairs understood as an increase (in agreement with most

doi: 10.3417/2009085 NOVON 21: 299–316. PUBLISHED ON 9 SEPTEMBER 2011.

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old classifications, e.g., Maximowicz, 1879; Zabel, TWO TAXONOMICALLY CONFUSED SPECIES OF SPIRAEA FROM

1893; Schneider, 1905; Rehder, 1927, 1940) or a NORTHERN YUNNAN

decrease (in agreement with the concept founded byTwo similar taxa of the genus Spiraea with simpleNakai, 1916; modified by Pojarkova, 1939, and Yu &

umbel-like, subsessile, lateral inflorescences on theKuan, 1963; and used in Chinese floras: Yu & Lu,previous year’s shoots and heteromorphic leaves were1974; Lu & Crinan, 2003; i.e., probably in agreementdescribed from northwestern Yunnan under thewith evolutionary trend). These morphotypes can benames S. calcicola W. W. Sm. and S. compsophylladivided and taxonomically applied in the followingHand.-Mazz. The former was described (Smith, 1913)way: (1) sessile flower fascicles (arising from lateralfrom an elevation of ca. 3500 m from Yulongxue Shanwinter buds) appearing as nonstalked umbels, with anorth-northwest of Lijiang. The latter name wasbasal leaf rosette; (2) shortly pedunculate (subses-described (Handel-Mazzetti, 1933) based on the

sile), congested corymbs, with a basal leaf rosette; (3)author’s own material from 3200 m in the mountains

corymbs or racemose (elongated) corymbs on asituated to the southeast above what is today Luoji at

peduncle that continues as a leafy short branchlet;the eastern margin of Zhongdian Xian near the

and (4) seeming umbels on a peduncle that continuesSichuan border. Although these localities are found

as a leafy short branchlet. The first and secondtoday in different prefectures (Lijiang and Diqing,

morphotypes have been traditionally classified as respectively), they are situated only 70 km from oneSpiraea sect. Glomeratae Nakai, the latter two as another. Yu and Kuan (1963) listed both taxaSpiraea sect. Chamaedryon Ser. in a narrower separately as members of Spiraea (sect. Glomeratae)concept (Yu & Kuan, 1963; Yu & Lu, 1974; ser. Hypericifoliae Pojark. ex T. T. Yu. Both taxa wereBusinsky & Businska, 2002). The term umbel-like also treated as separate species in Chinese floras (Yuor umbellate raceme used by several authors is & Lu, 1974; Lu & Crinan, 2003), and the sameequivocal and can be understood as a simple corymb, relevant illustration of both was given in one plate ini.e., an intermediate form between raceme and umbel both publications (Yu & Lu, 1974: 61, t. 8; Wu &(as used in the key and elsewhere in the Flora of Raven, 2004: 34). Beyond the various leaf shapesChina treatment by Lu & Crinan, 2003, in contrast to and fruits discussed for both taxa, the plate inumbel), or as the collective term for simple corymb question contained a flowering branchlet of S.and umbel (Rehder, 1927, 1940). Maxwell and Knees calcicola and a fruit-bearing branchlet of S. comp-(1995) used the term corymb for sessile flower sophylla, which are hardly comparable. Both taxafascicles and seeming umbels, or for the true (simple) were also accepted separately in the Index Floraeand also compound corymbs (they usually did not use Yunnanensis (Institutum Botanicum Kunmingensethe designation as compound). They alternatively also Academiae Sinicae, 1984: 542), with S. calcicolaused the term raceme, but only for S. media Schmidt. given from between 2700 and 3600 m elevation, andThe seemingly umbellate morphotype found in the S. compsophylla from between 2000 and 4000 m.genus Spiraea is only an approximate analogy, where Detailed study of the type material for both S.the pedicels are 6 equally long and densely crowded calcicola and S. compsophylla revealed both taxatogether, but they do not arise strictly from the same perfectly conspecific, characterized principally by thepoint. A few isolated flowers on a similarly long distinctly angled, dark brown shoots and branchlets.pedicel as shifted down on the peduncle may also By nomenclatural priority, Handel-Mazzetti’s name

often occur. The racemose corymb, as a variant of synonymizes to Smith’s older one.

simple corymbs within the above-mentioned thirdmorphotype, is an intermediate form between raceme Spiraea calcicola W. W. Sm., Notes Roy. Bot. Gard.

and corymb, and can be defined as an elongated Edinburgh 8: 131. 1913. TYPE: China.Yunnan: [Lijiang Xian], crevices of dry lime-corymb with the lower pedicels only indistinctlystone cliffs on the eastern flank of the Lichianglonger than upper ones. Racemose corymbs typicallyRange [Yulongxue Shan], 27occur, for instance, in

¡209N [27 N],S.¡10media. 9

11,000–12,000 ft., June 1910, G. Forrest 5730Together with my wife, Ludmila Businska, we(holotype, E not seen; isotype, PE). Figure 1.carried out a series of research expeditions to China,

where all taxa of the genus Spiraea that we found in Spiraea compsophylla Hand.-Mazz., Symb. Sin. 7: 450, pl.

nature were sampled (Businsky & Businska, 2002: 13, fig. 1. 1933, syn. nov. TYPE: China. Yunnan:‘‘Inter pagos Yungning und Dschungdien (Chungtien)11). In addition to these cited specimens, most[Zhongdian], in regionis temperatae rupibus aridis

Spiraea collections at PE and KUN were also montis Lamatso. Substr. Calceo, ca. 3200 m,’’ 12 Julyexamined. 1915, Handel-Mazzetti 7608 (holotype, W).

Shrubs to 1–1.5 m tall, densely branched, with pubescent to glabrescent, not cracking or peeling in thelonger branches 6 arcuate, angled, grayish dark brown following year; buds elongate-conical, usually shorterfrom second year; sterile long shoots of the present year or equal to half of petiole, with several sparsely pilose(fully developed only from July) to ca. 25( 40) cm, scales. Leaves deep green adaxially, grayish green andusually arcuate, dark reddish brown, conspicuously not papillose abaxially, with margin somewhat thick-angled with irregular ridges and grooves, sparsely ened, not revolute; venation reticulate, deep green,

Figure 1. Spiraea calcicola W. W. Sm. —A. Fertile branch with two young long shoots. —B. Three-year portion of sterilebranch with one young long shoot. —C. Inflorescence in fruit with basal leaf rosette. —D. Leaf from inflorescence basal rosette,abaxial view. —E, F. Fully developed leaves of sterile long shoots, abaxial view. —G. Fully developed leaf of sterile long shoot,adaxial view. Drawn by Ludmila Businska from July collection, R. Businsky 30409 (G), at the type locality.

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contrasting abaxially; leaf blades, as reduced in basal Flora of China treatment (Lu & Crinan, 2003: 72)part of sterile long shoots, often slightly longer than above all did not consider the sterile long shoots,wide, broadly angular-obovate and 6 narrowly cune- especially their specific leaf morphology. These areate; blades fully developed in middle and apical described only in Handel-Mazzetti’s Latin protologueportions of shoots, wider than long, transversally (1933) for the synonymous S. compsophylla based onrounded rhombic and broadly cuneate or truncate to the only specimen cited. A more complete descrip-subcordate at base, (5–)7–10( 12) 3 (6–)8–14( 16) tion and a new illustration are therefore given here,mm, 3-lobed or rarely 5-lobed, with notches shallow to based on additional specimens examined and from amore than half deep, all lobes 6 truncate or middle study of the species at two localities. However, bothlobe broadly triangular, in front irregularly crenulate, visits were during fruiting season, and thereforecrenate, or coarsely and obtusely dentate but with the flowers in blossom are not described, but expected toapical tooth often pointed; initially very sparsely pilose be in May or June.to almost glabrous 6 equally on both surfaces, more During our first expedition to Yunnan in 1990, wepilose only on base of principal veins abaxially; found and also collected Spiraea calcicola in thevenation consists of three principal veins branching region of its type locality on Yulongxue Shan, and theat one point at the triangular basal connection to second entity, represented to date by the name S.petiole; petioles (2–)3–5( 6) mm, 6 sparsely pilose, compsophylla, at its oldest documented locality nearwith basal lateral edges decurrent into longitudinal Qiaotou from 1939. We also found the former speciesshoot ridges, basal part persists after shedding the in 1998 at a new locality in the northeastern corner ofleaves; leaf blades of lateral rosettes at the inflores- Yunnan (near the Sichuan border) on Yao Shancence base and on lateral shortest sterile branchlets of (Qiaojia Xian).older twigs entire, obovate, with narrowly cuneate baseand rounded apex, (2.5–)4–7( 9) 3 (1.5–)2.5– Selected specimens. CHINA. Yunnan: Zhongdian Xian,3.5( 4.5) mm, glabrous on both surfaces, 3-veined; N flank of Haba Snow Range, K. M. Feng 1257 (KUN, PE);

Likiang [Lijiang] Xian, Likiang Snow Range, Zang 8938petiole to ca. 2 mm. Inflorescences densely arranged,(KUN, PE); Zhongdian Xian, Sanqi Xiang, ?Shen 21035

usually 10 to 30, set laterally on arcuate branches (KUN, PE); Lijiang Pref. & Xian, Yulongxue Shan, E foot(changed from previous year’s long shoots), each with above rd. from Lijiang to Daju, in valley ca. 2 km W ofbasal leaf rosette; basically shortly pedunculate Heishui, R. Businsky 30409 (G, Silva Tarouca Research

corymbs, on less vigorous branchlets congested, sessile Institute for Landscape and Ornamental Gardening [RI-LOG] herbarium); Zhaotong Pref., Qiaojia Xian, Yao Shan,and therefore of umbel-like form, on lower floweringupper NW slopes of great steep rocky amphitheatre above

portion of more vigorous branchlets with the peduncle Kangjiahe, R. Businsky 46405 (G, RILOG herbarium).to ca. 12 mm, with distant pedicels; corymbs with (6)7to 12(to 26) flowers on firm, glabrous, unbranched The following undescribed species is discriminat-

pedicel usually with narrow bract, rarely some lower ed here from material of the misinterpreted Spiraea

pedicels branched, with several flowers. Hypanthium compsophylla and is presented below as a new

campanulate, thickly walled, firm and fragile when dry, species.

externally entirely glabrous, smooth, light brown, insidewith usually sparse, long hairs, ca. 2.5 mm diam. in Spiraea adiantoides Businsky, sp. nov. TYPE:

fruit; sepals triangular, 0.7–1.431–1.6 mm, externally China. Yunnan: Diqing Pref., Zhongdian Xian,

glabrous, inside finely pubescent or glabrescent, erect- S foot of Habaxue Shan, Hutiaoxia (Tiger Leapspreading in fruit. Follicles erect-spreading to erect, Gorge), Jinsha Jiang (Yangtze river) valley ca. 4sparsely hairy only along ventral suture or entirely km ENE from mouth of Xiaozhongdian He atglabrous, with erect-spreading styles. Hutiaoxiazhen (Qiaotou), fissures of almost

vertical rocks of crystallic limestone, 1950 m,Habitat and distribution. Spiraea calcicola oc- 27¡119N, 100¡069E, 12 July 1990, R. Businsky

curs on rather sunny limestone, rocky, or scree 30403 (holotype, G; isotypes, MO, RILOGsubstrates between 2700 and 4000 m, often under herbarium). Figure 2.sparse pine stands of Pinus densata Mast. or P.armandii Franch. This taxon is known only from Haec species inter congeneros sinenses ad Spiraeamnorthwestern and northeastern Yunnan, China. sect. Glomeratas Nakai pertinentes quoad ramos elongatos

sicut ramulos teretes primo pallide brunneos atque foliaIUCN Red List category. Least Concern (LC; heteromorpha utrinque glabra Spiraeae kwangsiensi T. T.

IUCN, 2001). Yu simillima, sed ab ea ramis elongatis omnino glabrisatque ramorum elongatorum sterilium laminis foliaribus late

Notes. The protologue of Spiraea calcicola angulari-obovatis usque transverse rotundato-rhombicis,(Smith, 1913) and its subsequent description in the plerumque 3-lobis ac 3-veniis, abaxialiter manifeste

Figure 2. Spiraea adiantoides Businsky. —A. Fertile branch in July of second year. —B. Two sterile long shoots withsecondary lateral summer shoots. —C. Portion of fertile branch. —D. Inflorescence in fruit with basal leaf rosette. —E. Fullydeveloped leaf of sterile long shoot, abaxial view. Drawn by Ludmila Businska, parts A and E from the holotype, R. Businsky30403 (G), B–D from isotypes.

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cinereo-pruinosis venatione atroviridi discrepante distin- narrowly ligulate bract at base, the top pedicel withguitur. minute bract shifted up. Hypanthium campanulate,

thinly walled, externally usually entirely glabrousShrubs to 50 cm tall, pruinose and generally(sometimes with sporadic hairs), grayish brown,glabrous, with slender, straight, terete branches; longinside with long white hairs, 1.6–2.3 mm diam. inshoots and branchlets conspicuously slender, terete,fruit; sepals rounded triangular, 0.9–1.2( 1.4) mm,entirely glabrous, pale brown, initially smooth,externally glabrous, inside finely pubescent orcracking later, and peeling into longitudinal stripsglabrescent, erect to spreading in fruit; petals obovatein the second year; buds small, subconical, shorterwith retuse apex, 2.9–3.53 1.9–2.2 mm; stamens ca.than petiole, with several scales sparsely pilose20, shorter than petals, with filaments (1.3–)1.5–especially along margin, often sprouting during the1.7( 1.9) staminalfirst summer vigorous green mm; disk indistinct, interrupted,in shoots. Leaves paleformed by 10 small, rounded, lobed glandules;adaxially, conspicuously grayish pruinose and papil-carpels inside, including bases of styles, with longlose abaxially, with revolute margin; venationwhitish hairs. Follicles erect and parallel in maturity,reticulate, deep green, contrasting abaxially, in2.2–2.7 mm, sparsely hairy only along ventral suture;principal veins distinct adaxially; leaves entirelystyles ca. 1 mm, dorsal, almost spreading.glabrous on both surfaces or only some young leaves

with a few hairs on the petiole or principal veins Habitat and distribution. The new species occursabaxially; leaf blades of long shoots and lateral in fissures of open, sunny limestone rocks betweenrosettes or shortest branchlets unequal in shape and 1700 and 2200 m along the foot of high mountainsize; blades on sterile long shoots more variable, ranges, and is known only from two areas inbroadly angular-obovate to transversally rounded northwestern Yunnan, China, at a distance of ca.rhombic, often wider than long with broadly cuneate 150 km.to almost truncate base, or blades narrower, but at

IUCN Red List category. Vulnerable (VU; IUCN,most 1.53 longer than wide with a minimum base2001).angle ca. 55¡, (3–)6–12( 17)3 (2.5–)6–12( 18) mm,

excluding petiole; irregularly shallowly and orbicu- Phenology. Spiraea adiantoides flowers mostly inlarly 3-lobed or only crenate to almost entire, middle April to May.lobe truncate, retuse, or crenulate, lateral lobessimilar, but more irregular; venation consists of 3 Paratypes. CHINA. Yunnan (NW): Chungtien [Zhong-

principal veins branching at almost one point above dian], betw. Chiao-tou [Qiaotou] to Ya-chang-gou SE ofHaba Snow Range on Yangtze, 23 Mar. 1939, K. M. Fengbase; petioles 1–2.5 mm, with 1/3–1/2 remaining567 (KUN [2], PE); Gongshan, Sigitong, 2000–2200 m, 20

after leaf fall; leaf blades of lateral rosettes at the Sep. 1940, K. M. Feng 7988 (KUN [2], PE [3]); Gongshan,inflorescence base small, entire, obovate or spatulate, Azhong Luo, 1700 m, 25 May 1960, Chinese Collector 8746with narrowly cuneate base and rounded to obtuse (KUN, PE [2]).

apex, to 6.2 3 2.7 mm, including the petiole to 0.5mm long, with one principal vein pinnately branched; TWO TAXONOMICALLY CONFUSED SPECIES OF SPIRAEA FROMblades on lateral shortest sterile branchlets of older TAIWAN

twigs (usually also in rosettes) similar but somewhatmore variable, to 9 3 4 mm including petiole, entire According to relevant Chinese floras (Li, 1963; Yuor often shallowly 3- or 4-lobed; leaves on long shoots & Lu, 1974; Ohashi & Hsieh, 1993; Lu & Crinan,persist usually until second half of following year, 2003), only five species of the genus Spiraea arethus inflorescences with the basal rosette of small known from Taiwan. Three of them, S. formosanaentire leaves appear in axils of larger, lobed, broadly Hayata, S. hayatana H. L. Li, and S. morrisonicolacuneate leaves from previous year. Inflorescences Hayata, all with terminal compound corymbs, areumbel-like, congested, few-flowered corymbs set traditionally classified into section Calospira K.laterally on straight branchlets (changed from Koch. Of the two species with simple inflorescences,previous year’s long shoots); corymbs sessile or more S. tarokoensis Hayata from section Chamaedryon hasoften on (1–)2–3( 4) mm peduncle with basal leaf umbel-like corymbs on short, lateral, leafy branchletsrosette in axil of previous year’s leaf or its remnant on the previous year’s shoots, and S. prunifoliapetiole, bearing (3)4 or 5(to 7) flowers, each always Siebold & Zucc. from section Glomeratae has sessilesingly on pedicel 5–7( 9) mm, slender, lax, usually flower fascicles on the previous year’s shoots (bothentirely glabrous (sometimes with only sporadic according to the infrageneric classification of Yu &hairs), grayish, the lower pedicel usually with Kuan, 1963). In November 2008 during the ascent ofdiminished leaf at base, upper pedicels with small, the highest Taiwanese mountain, Yushan (Mt. Jade,

Volume 21, Number 3 Businsky 3052011 Review of Chinese Spiraea (Rosaceae)

3952 m), along the classic tourist path from Tataka pinnate with usually 3 pairs of lateral veins leading toVisitor Center, the present author found a sporadic, teeth apex, midvein and lateral veins raised abax-low shrub of Spiraea growing on a sparsely wooded ially; petioles 2–3 mm, with dense, long, sericeousslope at 2800 m elevation (cf. R. Businsky 66401). hairs. Inflorescences seeming umbels on lateral,This shrub represents the group of typically seem- slender, leafy, 1–4 cm branchlets, with ca. 10 to 15ingly umbellate species hitherto classified as an flowers, each always singly on slender, villous, ca. 1unspecified part of series Trilobatae Pojark. ex T. T. cm pedicel. Hypanthium pubescent on both surfaces;Yu from Spiraea sect. Chamaedryon, i.e., the group of sepals triangular, ca. 1 mm, externally glabrous,several mostly Chinese species not known from inside pubescent, spreading; petals white, suborbic-Taiwan according to the above-mentioned Chinese ular, 2–3 3 ca. 3 mm, slightly emarginate; stamensfloras. However, recently, during the compilation of a ca. 20, shorter than petals, with filaments ca. 1.5–2Red Data Book for endangered and threatened plants mm; carpels with long hairs inside and distally, styleof Taiwan, one previously published species, S. glabrous, ca. 1.5 mm. Follicles initially erect, ca. 2.5tatakaensis I. S. Chen, was unexpectedly found mm, sparsely hairy chiefly along ventral suture neardescribed and illustrated thoroughly in a Chinese style insertion.journal (Chen, 1978) not familiar to the taxonomists.

Habitat and distribution. Spiraea tatakaensis oc-Thus this taxon has been overlooked by previouscurs on sunny forest margins or sparse woody slopes onbotanists and could not be found in any works on theslate or silicate ground, mostly between 1200 and 3200flora of Taiwan or China. The careful morphologicalm, often among trees of Pinus taiwanensis Hayata.comparison reveals that the above-mentioned shrubKnown only from one area in the central mountainfound by the present author is conspecific with S.range of Taiwan, particularly around the highest massiftatakaensis. Because the protologue of this species isYushan (Mt. Jade, 3952 m), S. tatakaensis isnot easily available, the accurate description (com-documented from the western and northeastern sidespiled from R. Businsky 66401 and the originalnear the border of Chiayi and Nantou counties, anddescription and illustration) and the new illustrationfrom a locality in Tanta valley ca. 35 km north-are given here.northeast of Mt. Yushan, in Nantou County.

Spiraea tatakaensis I. S. Chen, Formosan Sci. 32(3): IUCN Red List category. Least Concern (LC;53. 1978. TYPE: Taiwan. Tataka, Mt. Yuishan, IUCN, 2001).19 June 1977, I. S. Chen 5129 (holotype, TAIFnot seen) Figure 3. Phenology. Spiraea tatakaensis flowers mostly in

May to July.Shrubs to ca. 1 m tall, with relatively slender,

terete, straight branches and shoots; long shoots Notes. The five cited collections of Spiraea

densely grayish tomentose initially, beginning crack- tatakaensis at TAI and TAIF were incorrectly

ing and peeling in lower mature part at end of the first determined as S. tarokoensis and were also misiden-

year; buds ovoid to obtusely conical, 1–2 mm, shorter tified in the appendix to the new Flora of Taiwan

than petiole, with several scales pilose on exterior treatment (Ohashi & Hsieh, 1993: photo 12), listing

surface initially, at least along margin later. Leaves the former species under the name of the latter.coriaceous; deep green, finely rugose, with immersed Spiraea tarokoensis is locally endemic, known onlyveins, and sparsely sericeous adaxially; glaucous, from one limited area above the eastern coast ofdistinctly papillose, with fine, contrasting venation, Taiwan, where it occurs at relatively low elevations.and laxly sericeous abaxially, with hairs more dense This rare species is inexactly described in theon prominent principal veins and on revolute margin; literature. According to Yu and Kuan (1963) andleaf blades angular-obovate to narrowly obovate, 1.5– Yu and Lu (1974), S. tarokoensis is placed within the3.53 longer than wide, narrowly cuneate proximally series Mediae Pojark. ex T. T. Yu, vaguelybut often obtuse at very base, (15–)20–30( 35)3 7– circumscribed from series Trilobatae (in which S.17 mm; margin with 3 or 4 pairs of decreasing, tatakaensis should be placed), based chiefly on theincised, forward-pointed, coarse, acute teeth in upper character of stamens shorter or subequal to petalshalf on sterile long shoots or usually with only 2 tooth versus stamens longer or slightly shorter than petalspairs in upper third of narrower leaves on short fertile in the former. Although S. tarokoensis can bebranchlets or in lower part of sterile long shoots; classified into the same group of seemingly umbellatelower pair of largest teeth in broader leaves often in species as S. tatakaensis, both species are quiteform of indistinct lateral lobes with 1 to 3 outer, dissimilar to each other, morphologically and alsominute, serrate teeth; apex of blade acute; venation ecologically; S. tatakaensis is compared to S.

Figure 3. Spiraea tatakaensis I. S. Chen. —A. Branch in autumn of second year with fertile short branchlets in the terminalpart after pedicels fell off. —B. Sterile long shoot. —C. Fully developed leaf of sterile long shoot, abaxial view. —D. Shortbranchlet with inflorescence. Drawn by Ludmila Businska, parts A–C from November collection, R. Businsky 66401 (G), D fromSheng-You Lu 14766 (TAIF), both from the type locality.

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tarokoensis and other seemingly umbellate species in upward, leading to teeth apex, raised abaxially;Table 1. There are several discrepancies concerning petioles 1.5–5 mm, villous; leaf blades on shortS. tarokoensis in the Flora of China treatment (Lu & fertile branchlets obovate, elliptic, or ovate, 7–16Crinan, 2003) and in Taiwanese floras (Li, 1963; Liu mm, crenate or crenate to 3-lobed in upper third.& Su, 1977; Ohashi & Hsieh, 1993). The illustration Inflorescences seeming umbels on lateral, slender,in the new Flora of Taiwan treatment (Ohashi & leafy, 0.5–2.5( 5) cm, villous-tomentose branchlets,Hsieh, 1993: 156, pl. 71) is ascribed to S. with 8 to 20 flowers, each always singly on slender,tarokoensis, but is clearly not this species. Rather, villous, 4–12 mm pedicel. Hypanthium and sepalsplate 71 is certainly a drawing of S. prunifolia, villous on both surfaces; hypanthium shallowlyapproximately corresponding to S. prunifolia var. funnel-shaped; sepals triangular with acute apex,pseudoprunifolia (Hayata ex Nakai) H. L. Li. There is 0.7–1.3 mm, spreading to reflexed; petals trans-a second drawing of allegedly the latter taxon (but versally obovate, emarginate, ca. 2 3 3 mm; stamensillustrated incorrectly with pubescent follicles) on the shorter than petals. Follicles suberect, 1.7–2.4 mm,preceding plate 70 (Ohashi & Hsieh, 1993: 153). with long hairs particularly on dorsal side; styles ca.Given this confusion, an accurate description and the 0.7–1.3 mm, dorsal, erect-spreading.first illustration of S. tarokoensis correctly ascribed

Habitat and distribution. Spiraea tarokoensisare given here, with both based on our collectedoccurs on open limestone rocks or rocky sitesmaterial and the type.between 400 and 1300 m. It is known only from

Selected specimens. TAIWAN. Nantou Co.: Tong Puh– the limited limestone area around Tienhsiang aboveMt. Morrison [Yushan], Ih-Sheng Chen 3750 (TAI); Taroko gorge on the Pacific side of Taiwan’s centralSinyihsian, Dandalindao [Tanta valley], Aleck T. Y. Yang mountain range, northwest of Hualien.et al. 6038 (TAI); Loloku–Kuankao, S. F. Huang 5273(TAI). Chiayi Co.: [betw.] Tatachia [Tataka] & Paiyun IUCN Red List category. Spiraea tarokoensis isShelter, Sheng-You Lu 14766 (TAIF); en route to Yushan, certainly one of the rarest Spiraea species in theT. C. Huang et al. 14253 (TAI); Yushan Natl. Park, Yushan

world. The difficult accessibility of rocky sites(Mt. Jade) massif, S slopes of Mt. Jade Front Peak (3239 m),along path from Tataka (Paiyun) visitor center to Paiyun harboring the taxon is the most favorable factor inLodge, R. Businsky 66401 [Spiraea abscondita Businsky, protecting it. However, a real risk for this species cannom. inval., in sched.] (G, RILOG herbarium). be caused by wildfires or by construction or widening

of roads (the population sampled by the presentauthor at 950 m in 1991 was not found in 2008 due toSpiraea tarokoensis Hayata, Icon. Pl. Formosan. 9:road erosion and built-up protection). Therefore, this38–39. [25 Mar.] 1920. TYPE: Taiwan. Hualien,taxon is considered Vulnerable (VU) according toTaroko, 27 Apr. 1917, B. Hayata & S. SasakiIUCN Red List criteria (IUCN, 2001).s.n. (types, TI [2]). Figure 4.

Phenology. This species has been collected inLax shrubs to ca. 1 m tall, with relatively slenderflower in April to May.branches; shoots basically terete, with slender

longitudinal ridges and soon cracking and peelingSelected specimens. TAIWAN.surface, villous-tomentose initially; buds globose to Hualien Co.: Nai-taroko

[Taroko Natl. Park], Kwarenko, B. Hayata & S. Sasaki 68ovoid, with many keeled, villous scales, often (TAIF 12535, 12536 [s.n.], 12537); [Taroko Natl. Park]sprouting in vigorous long shoots and also in fertile Yenhai logging trail, Shih-Wen Chung 8006 (TAIF), Pi-branchlets into lateral summer shoots (usually longer Fong Lu 9695 (TAIF); Taroko Natl. Park, limestone rocks of

than bearing fertile branchlets). Leaves pale green, NE exposure along Central Cross-Island Hwy. near viewabove Tienhsiang (Tiansiang) ca. 1 km E of Hsipo hamlet,almost glabrous and with not immersed veinsR. Businsky 32407 (G, RILOG herbarium).

adaxially, almost glabrous, glaucous and papilloseabaxially, with revolute margin; venation reticulate,contrasting and on principal veins pubescent abax- TAXONOMIC CHANGES IN SPIRAEA FOR FLORA OF CHINA

ially; leaf blades on sterile long shoots usually ovateto suborbicular, sometimes rhombic-ovate, 1.2–23 Three taxa accepted in the Flora of Chinalonger than wide, with rounded or broadly rounded treatment (Lu & Crinan, 2003) as independentcuneate base, (8–)12–253 (7–)10–20 mm, shallowly species are taxonomically changed here, either3-lobed or coarsely crenate mostly only in upper half, replaced in Chinese floras by a neglected taxon withwith lobes or teeth 6 obtuse, usually secondarily a newly distinguished variety, synonymized, orcrenate, and with obtuse to rounded apex; venation relegated to the lower rank. Several new synonymsconsists of 3 principal veins sparsely branched are also given.

1. Spiraea lasiocarpa Kar. & Kir., Bull. Soc. Imp. [43¡099N, 81¡079E], s.d., Xinjiang exped. 10530

Naturalistes Moscou 15: 536. 1842. TYPE: (holotype, PE).

[Kazakhstan.] ‘‘Dzhungaria, ad fl. Sarkan,’’ s.d., Notes. The names Spiraea mongolica KoehneG. S. Karelin 1441 (holotype, LE). (Koehne, 1893: 212) and S. gemmata Zabel (Zabel,

1893: 23) are referred to the autonymic variety of S.1a. Spiraea lasiocarpa var. lasiocarpa.lasiocarpa. Both Koehne’s and Zabel’s names require

Spiraea mongolica Maxim., Bull. Acad. Imp. Sci. St.- nomenclatural investigation.Petersbourg 27: 467. 1881. Basionym: Spiraea cren-ifolia C. A. Mey. c mongolica Maxim., Acta Horti

var. Businsky, var.Petropol. 6: 181. 1879. TYPE: [China.] ‘‘Mongolia 1b. Spiraea lasiocarpa villosa

occid., Terra Ordos [western Nei Mongol], montibus nov. TYPE: China. Sichuan: Garze Pref., DawuAlaschan [Helan Shan], pars media declivit. occid.,’’ 18 Xian, Dawu, foot of steep NE slopes above rightJune 1873, Przewalski 295 (holotype, LE; isotype, PE). bank of Xianshui He river SW opposite to town,

Spiraea mongolica Maxim. var. tomentulosa T. T. Yu, Acta grassy slope on limestone ground, 3100 m,Phytotax. Sin. 8(3): 216. 1963, syn. nov. TYPE:

30¡599N, 101¡079E, 1 June 1992, R. BusinskyChina. [Nei Mongol:] ‘‘Mongolia Interiora, Mt. Ala, inrupibus siccis, 1500–1700 m,’’ Aug. 1933, W. Y. Hsia 33403 (holotype, G; isotype, RILOG herbarium).3893 (holotype, PE).

Spiraea mongolica Maxim. var. pubescens Y. Z. Zhao & T. J. A varietate typica ramis novellis gemmis pedunculis etWang, Bull. Bot. Res., Harbin 20(3): 259. 2000, syn. pedicellis villoso-tomentosis, foliis utrinque aequaliternov. TYPE: China. Xinjiang: [Ili Pref.], Zhaosu villosis, hypanthiis et item sepalis extus villosis differt.

Table 1. Comparison of Spiraea tatakaensis and S. tarokoensis from Taiwan with similar seemingly umbellate species occurringin mainland China.

Character S. tatakaensis S. tarokoensis S. siccanea S. nishimurae

Shoot indumentum densely grayish tomentose villous-tomentose pubescent pubescent(initial)

Leaf blade surfaces deep green adaxially; pale green adaxially; deep green adaxially; light green on bothglaucous, distinctly almost glabrous and almost glabrous or surfaces; sparselypapillose, with only on principal only on principal pubescent andcontrasting venation, veins pubescent, veins pubescent and without contrastingand laxly sericeous glaucous and papillose gray-green abaxially venation andabaxially abaxially papillae abaxially

Leaf blades on sterile long shootsLength, cm mostly 2–3 mostly 1–2.5 1–2.5 mostly 1–2Shape angular-obovate to ovate to suborbicular, obovate or oblong mostly rhombic-ovate

narrowly obovate, 1.5– 1.2–23 longer than to elliptic3.53 longer than wide wide

Base narrowly cuneate rounded or broadly broadly cuneate to narrowly cuneaterounded cuneate subrounded

Margin with 3 or 4 pairs of shallowly 3-lobed or shallowly 3-lobed and with 2 or 3 pairs ofdecreasing, coarse, coarsely crenate in crenate above coarse, secondarilyacute teeth in upper 1/2 upper 1/2, with lobes middle serrate teeth or

or teeth 6 obtuse, lobes in upper 2/3secondarily crenate

PetiolesLength, mm 2–3 1.5–5 2–5 1–3Indumentum dense, long, sericeous villous sparsely pubescent or sparsely pubescent

hairs glabrousInflorescences ca. 10- to 15-flowered 8- to 20-flowered mostly 7- to 15- 7- to 25-flowered

flowered

PedicelsLength, mm ca. 10 4–12 7–10 6–10Indumentum villous villous densely pubescent sparsely pubescent

Distribution central Taiwan eastern Taiwan northwestern Yunnan north-central tonortheastern China

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This variety differs from Spiraea lasiocarpa var. and later known from floras of Pakistan (Stewart,lasiocarpa by the shoots and buds villous-tomentose 1972) and India (Purohit & Panigrahi, 1991). The(vs. mostly glabrous or pubescent in the autonymic autonymic variety representing the species was foundvariety), by the leaves including petioles 6 densely to be taxonomically identical with S. mongolicavillous equally on both blade surfaces including Maxim. (Businsky & Businska, 2002: 21), describedmargin (vs. blades entirely glabrous or puberulous from what is today the Chinese province of Neialong usually lower margin portions), by corymbs with Mongol, and as presented in Chinese floras (Yu & Lu,the peduncle and pedicels villous-tomentose (vs. 1974; Fu & Hong, 2003; Lu & Crinan, 2003).entirely glabrous), and by the hypanthium and sepals Spiraea gemmata Zabel, another taxon conspecificvillous externally (vs. glabrous). with S. lasiocarpa, now to its autonymic variety, was

described from cultivated plants of northern ChineseNotes. Spiraea lasiocarpa was described from the´ ´Dzhungarskij Alatau Mountains (today’s eastern origin (Businsky & Businska, 2002: 28), and this

Kazakhstan) from a locality close to the present name has previously been used often in the literature

Chinese border. It has been missing in important (Schneider, 1905; Rehder, 1913, 1927, 1940; Bean,

dendrological or specialized works (Zabel, 1893; 1981; Maxwell & Knees, 1995) for the relevantSchneider, 1905; Rehder, 1927, 1940; Bean, 1981), Chinese plants. The correct name for this speciesor in Chinese floras (Yu & Lu, 1974; Lu & Crinan, widely distributed from Central Asia and the2003), but known from Russian literature (e.g., northwestern Himalayas to north-central China is S.Pojarkova, 1939; Shul’gina, 1954; Svjazeva, 1980) lasiocarpa, by its nomenclatural priority. None of the

Table 1. Extended.

S. pubescens S. hirsuta S. dasyantha S. yunnanensis

pubescent pubescent densely grayish tomentose densely grayish tomentose

pubescent on both surfaces, deep green adaxially; only deep green adaxially; deep green adaxially;not papillose abaxially sparsely pubescent densely whitish tomentose densely gray tomentose

abaxially intially abaxially abaxially

2–6 2.5–6.5 1.5–4.5 1–3.5rhombic-ovate to elliptic obovate, rhombic-ovate, or rhombic-ovate, 1.1–1.83 ovate, suborbicular, or

elliptic longer than wide obovate

narrowly cuneate or rounded narrowly to broadly cuneate broadly cuneate to broadly cuneate, roundedcuneate or rounded cuneate subcordate to subcordate

with irregular coarse, serrate or serrate-lobed in with 3 to 5 pairs of shallowly crenate tosometimes secondarily upper 1/2 or 2/3 secondarily serrate teeth incised obtusely lobedserrate teeth, or shallow or lobes except base except base, secondarilylobes in upper 1/2 or 2/3 serrate

2–6 5–10 2–5 1–6pubescent pubescent densely tomentose densely tomentose

15- to 40-flowered . 20-flowered 10- to 25-flowered 5- to 30-flowered

5–15 mostly 10–20 6–10 3–10glabrous or glabrescent densely pubescent densely gray tomentose densely gray tomentosecentral to northeastern central to southeastern China central to eastern China Yunnan and westernChina; Mongolia, Russia, SichuanKorea

Volume 21, Number 3 Businsky 3092011 Review of Chinese Spiraea (Rosaceae)

Figure 4. Spiraea tarokoensis Hayata. —A. Sterile long shoot with secondary lateral summer shoots. —B. Fertile branch withthree-year basal portion and summer shoots grown laterally from bearing short fertile branchlets. —C. Fully developed leaf ofsterile long shoot, abaxial view. —D. Short branchlet with inflorescence. —E. Fully developed leaf of sterile long shoot, adaxialview. Drawn by Ludmila Businska, parts A–C from December collection, R. Businsky 32407 (G), at the type locality; D and Efrom the type, B. Hayata & S. Sasaki s.n. (TI).

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mentioned taxa are reported from the neighboring Spiraea oblongifolia Waldst. & Kit., Descr. Icon. Pl. Hung.Mongolia (Grubov, 1982). The indumentum of S. 3: 261, t. 235. 1812. TYPE: [Hungary.] ‘‘Spiraealasiocarpa (including the follicles) varies from oblongifolia Metsekensis ex horto,’’ s.d., P. Kitaibelentirely glabrous morphotypes to wholly villous ones. s.n. (lectotype, designated by Kovats [1992: 50], BP).

Whereas the typical form is close to the first andNotes. The name Spiraea confusa Regel & Korn,glabrous extreme, the second one can be accepted at

published in 1857, has been used in the olderthe rank of variety as newly described below. Theliterature (e.g., Rehder, 1927; Pojarkova, 1939), buttaxa described as S. mongolica var. tomentulosa andno type was designated. The name is referred to theS. mongolica var. pubescens represent not markedautonymic variety of S. media.forms within the range of indumentum variation of S.

lasiocarpa and are more closely affined to the typicalform, therefore they are accepted as new synonyms 3b. Spiraea media var. sericea (Turcz.) Maxim.,

for variety lasiocarpa. The opinion that S. mongolica Acta Hort. Petrop. 6: 189. 1879. Basionym:

var. tomentulosa and S. ningshiaensis T. T. Yu & L. Spiraea sericea Turcz., Bull. Soc. Imp. Natural-T. Lu are conspecific (Zhao & Wang, 2000) is neither istes Moscou 16: 591. 1843. TYPE: [Russiaaccepted in the Flora of China treatment (Lu & (Chita Province) & China (NE Nei Mongol)Crinan, 2003), nor here, because these taxa have a border.] ‘‘Dahuria, in lapidosis ad fluviumwholly different morphology in their winter buds (cf. Argun,’’ s.d., N. S. Turczaninow s.n. (type, LE).the following key).

Notes. Spiraea sericea was accepted in Russianliterature (e.g., Pojarkova, 1939; Shul’gina, 1954;2. Spiraea muliensis T. T. Yu & L. T. Lu, ActaSvjazeva, 1980) and in Chinese floras (Yu & Lu,Phytotax. Sin. 13(1): 101. 1975. TYPE: China.

Sichuan: Muli, Mengdong Shan, 2700 m, 21 1974; Lu & Crinan, 2003; Fu & Hong, 2003); it

June 1960, Ying Jun-Sheng 4102 (holotype, PE). differs from typical S. media only by leaf blades laxlysericeous abaxially and initially pilose adaxially (vs.

Spiraea daochengensis L. T. Lu, Bull. Bot. Res., Harbin blades glabrous or almost glabrous in S. media var.9(2): 49. 1989, syn. nov. TYPE: China. Sichuan:Daocheng, 3800 m, 1 Sep. 1981, Qing-Zang Exped. media). Because the former taxon falls within the5947 (holotype, PE). range of morphological variation and also geograph-

ical distribution of the latter, the former is acceptedNotes. These two imperfectly known taxa from

here only in the rank of variety.southwestern Sichuan were accepted as separatespecies in the Flora of China treatment (Lu & Crinan,2003). However, they are now found to be conspecific KEY TO THE CHINESE SPECIES OF SPIRAEA

after careful comparison of both holotypes, as well asThe following key contains all Chinese speciesbased on a study of the species variation at the newly

accepted by the author for the genus Spiraea withfound locality specified below. The holotype ofSpiraea daochengensis represents a form with sparser lateral, simple inflorescences (with 1-flowered pedi-

indumentum on shoots, buds, both leaf blade cels), usually classified into the two sections, Spiraea

surfaces, pedicels, and hypanthia including sepals sect. Glomeratae and Spiraea sect. Chamaedryon.

externally, whereas the holotype of S. muliensis has This taxonomic key substitutes for the correspondingall mentioned element surfaces densely villous. portion in the Flora of China treatment (Lu & Crinan,

2003: 49–50, key leads 30–70), where severalSpecimen collected. CHINA. Sichuan: Garze Pref., discrepancies were found. The crucial diagnostic

Jiulong Xian, bottom of upper Taka He valley, 3300 m,¡ ¡ character of the inflorescences as compound versus

29 019N, 101 499E, 5 July 2001, R. Businsky 51405 (G,RILOG herbarium). simple, is missing in the lead couplet (1) in the Flora

of China key. Alternative constructions of some¨3. Spiraea media Schmidt, Osterr. Allg. Baumz. 1: leads, believed more useful for determination, were

53, t. 54. 1792. TYPE: [Russia, West Siberia.] compiled for the key presented here, which also‘‘Sibiria occid., prope fl. Irtys,’’ 1733–1736, J. excludes other less reliable characters previouslyG. Gmelin s.n. (type, LE not seen).

used. Significant synonyms are given in parenthesesin the key. Altogether, 40 species of Spiraea and one

3a. Spiraea media var. media. interspecific hybrid are included in the following key.

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KEY TO THE CHINESE SPECIES OF SPIRAEA WITH SIMPLE INFLORESCENCES

1a. Inflorescences terminal or lateral, always compound, with branched pedicels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [28 Chinese species outside this study]

1b. Inflorescences always lateral, simple, with unbranched, 1-flowered pedicels (rarely some lower pedicels poorlybranched).2a. Flowers in sessile fascicles or shortly pedunculate (subsessile) crowded together corymbs, only with basal

leaf rosette (Spiraea sect. Glomeratae).3a. Leaves heteromorphic, those on sterile long shoots usually distinctly lobed or at least coarsely crenate-

serrate, those in basal rosettes under inflorescences not lobed, entire.4a. Long shoots and branchlets conspicuously angled, dark reddish brown initially . . . S. calcicola W. W. Sm.4b. Long shoots and branchlets terete, pale brown initially.

5a. Leaf blades glabrous on both surfaces.6a. Long shoots entirely glabrous; leaf blades on sterile long shoots broadly angular-obovate to

transversally rounded rhombic, mostly 3-lobed, 3-veined, conspicuously grayish pruinose,with contrasting venation abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. adiantoides

6b. Long shoots pubescent or puberulent initially; leaf blades on sterile long shoots obovate,elliptic, or suborbicular, mostly not lobed, usually pinnately veined, pale green orglaucous green and usually without contrasting venation abaxially . . . . S. kwangsiensis T. T. Yu

5b. Leaf blades densely pubescent or tomentose abaxially.7a. Leaf blades on sterile long shoots mostly , 12 mm, 6 transversally obovate, flabellate, 6

deeply 3- to 5-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aquilegiifolia Pall.7b. Leaf blades on sterile long shoots mostly . 12 mm, broadly ovate, obovate, or elliptic to

suborbicular, coarsely crenate or shallowly 3-lobed, often secondarily crenate.8a. Shoots tomentose; leaf blade often 3-lobed, with a few coarse, obtuse teeth above

middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. martini H. Lev.8b. Shoots pubescent; leaf blade not lobed, with many coarse, crenate teeth above base

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cavaleriei H. Lev.3b. Leaves homomorphic, those on sterile long shoots similar in shape to those in basal rosettes below

inflorescences (disregarding numbers in serration or teeth).9a. Leaf blades entire or at most with 2 to 4 coarse teeth near apex; long shoots and branchlets terete.

10a. Shoots glabrous or puberulent initially; leaf blades usually 15–20 mm; follicles glabrous. .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hypericifolia L.

10b. Shoots 6 densely pubescent initially; leaf blades shorter than 15 mm; follicles pubescent. . . . . . . . . . . . . . . . . . . . . . . . S. chailarensis Liou (including S. arenaria Y. Z. Zhao & T. J. Wang)

9b. Leaf blades sharply serrate at least above middle (when only near apex then serration minute); longshoots and branchlets 6 angled.11a. Leaf blades linear-lanceolate, glabrous on both surfaces . . . . . . . S. thunbergii Siebold ex Blume11b. Leaf blades oblong, elliptic, or ovate, usually pubescent or villous on both or at least one

surface.12a. Leaf blades minutely serrate; long shoots and branchlets slightly angled; pedicels . 10

mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. prunifolia12b. Leaf blades coarsely serrate; long shoots and branchlets conspicuously angled; pedicels

, 10 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. muliensis2b. Flowers in corymbs, racemose corymbs, or seeming umbels, all on a peduncle continuing to leafy short

branchlet (Spiraea sect. Chamaedryon, s. str.).13a. Winter buds with 2 exterior valvate scales with acuminate apex (cf. Note 1).

14a. Leaf blades sharply serrate; stamens longer than petals; sepals reflexed in fruit.15a. Leaf blades lanceolate, oblong-elliptic, or oblong-ovate, margin simply serrate above

middle or only near apex; stamens ca. 20 . . . . . . . . . . . . . . . . . . S. flexuosa Fisch. ex Cambess.15b. Leaf blades broadly ovate, margin usually doubly serrate almost from base or above middle;

stamens 35 to 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chamaedryfolia L.14b. Leaf blades entire or apically crenate-serrate; stamens nearly equaling petals; sepals erect or

spreading in fruit.16a. Sterile shoots relatively long and slender; buds usually 2–5 mm; branchlets usually

straight; leaf blades on sterile long shoots usually oblong-elliptic . . . . . . . . . . . . . S. lasiocarpa16b. Sterile shoots relatively short and thick; buds usually 5–8 mm; branchlets arcuate; leaf

blades on sterile long shoots ovate, elliptic, or obovate, rather broadly so . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. mollifolia Rehder (cf. Note 2)

13b. Winter buds with several exterior 6 imbricate scales usually with obtuse or acute apex.17a. Prostrate shrub; flowers in 6 congested corymbs on a peduncle continuing to a very short

branchlet with leaves almost crowded to basal rosette; (leaf blades broadly ovate, 0.8–1.53 0.5–1 cm, long sericeous on both surfaces) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pjassetzkii Buzunova (including S. prostrata Maxim., nom. illeg.)

17b. Erect shrub, although sometimes lax; short branchlets bearing inflorescence with leaves arrangedat 6 distinct intervals.

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18a. Leaf blades usually , 8 mm wide and 2.5–53 longer than wide, entire, or sometimes with afew teeth near apex.19a. Long shoots and branchlets terete, glabrous; sepals reflexed . . . S. dahurica (Rupr.) Maxim.19b. Long shoots and branchlets angled, pubescent initially; sepals erect or almost spreading.

20a. Leaves glabrous on both surfaces; petals white; follicles with subterminal,divergent styles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alpina Pall.

20b. Leaves pubescent on both surfaces; petals pinkish to purple-red; follicles withdorsal, suberect styles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . S. xizangensis L. T. Lu (including S. tibetica T. T. Yu & L. T. Lu, nom. illeg.)

18b. Leaf blades on sterile long shoots usually . 8 mm wide and , 2.53 longer than wide,crenate, serrate, lobed, or sometimes entire.21a. Long shoots and branchlets angled.

22a. Leaf blades broadly ovate, often sparsely denticulate above middle, sometimesshallowly 3-lobed, 3-veined; inflorescences , 2 cm wide . . . . . . S. ningshiaensis

22b. Leaf blades elliptic, elliptic-oblong, or obovate-oblong, entire or sometimesdenticulate at apex, pinnately veined; inflorescences usually . 2 cm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. myrtilloides Rehder

21b. Long shoots and branchlets terete or almost terete.23a. Flowers in corymbs or racemose corymbs (pedicels 6 distant on inflorescence

axis); stamens at least slightly to ca. 23 longer than petals.24a. Leaf blades entire or coarsely serrate at apex or at most in upper third

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. media24b. Leaf blades simply or doubly, sometimes obtusely serrate, incised

serrate or serrate lobed, at least above middle.25a. Shoots, leaf blades abaxially, pedicels, hypanthia externally, and

follicles pubescent; sepals erect in fruit . . . S. sublobata Hand.-Mazz.25b. Shoots, leaf blades abaxially, pedicels, and hypanthia externally

usually glabrous or nearly so, follicles glabrous, sometimes pilosealong ventral suture or throughout (S. anomala Batalin).26a. Leaf blades oblong-elliptic, oblong-ovate, or lanceolate-

elliptic, with base cuneate; sepals erect in fruit. . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. elegans Pojark.

26b. Leaf blades ovate, triangular-ovate, elliptic-ovate, orbroadly ovate, when lanceolate-ovate (sometimes in S.anomala) then with base truncate or subrounded; sepalsusually reflexed or sometimes suberect in fruit.27a. Corymbs 3- to 6-flowered, flowers 6–9 mm diam.;

follicles hirsute throughout . . . . . . . . . . . . . . . . . S. anomala27b. Corymbs 5- to 20-flowered, flowers 8–12 mm diam.;

follicles glabrous or pilose along ventral suture.28a. Leaf blades on sterile long shoots usually

incised doubly serrate . . . . . . . . . . . S. laeta Rehder28b. Leaf blades on sterile long shoots usually

shallowly simply or doubly serrate.29a. Leaf blades sericeous-villous on both

surfaces, gray-green abaxially; inflores-cences 12- to 20-flowered; flowers 8–10mm diam. . . . . . . . . . . . . . S. papillosa Rehder

29b. Leaf blades glabrous on both surfaces,pale-green abaxially; inflorescences 5- to10-flowered; flowers 10–12 mm diam. .. . . . . . . . . . . . . . S. lichiangensis W. W. Sm.

23b. Flowers in seeming umbels (pedicels densely crowded almost into oneinsertion point at apex of inflorescence axis); stamens shorter than orsubequaling petals.30a. Leaf blades glabrous or almost glabrous on both surfaces or puberulous

on veins abaxially.31a. Shoots pubescent or villous-tomentose initially.

32a. Shoots villous-tomentose initially, leaf petioles villous(eastern Taiwan) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tarokoensis

32b. Shoots pubescent initially, leaf petioles sparsely pubescentor glabrous (northwestern Yunnan) . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . S. siccanea (W. W. Sm.) Rehder

31b. Shoots glabrous from initiation.33a. Leaf blades on sterile long shoots rhombic-lanceolate to

rhombic-oblong, with base narrowly cuneate and apex acute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cantoniensis Lour.

314 Novon

33b. Leaf blades on sterile long shoots rhombic-ovate, rhombic-obovate, broadly ovate, or suborbicular, with base 6 broadlycuneate (sometimes shortly narrowed toward petiole),rounded or subcordate, and apex obtuse or subacute.34a. Leaf blades bluish gray abaxially; fruits often not

developing or irregularly developing (cultivatedplants of cultural origin) . . . . . . . . . S. 3vanhouttei (Briot)

Carriere (¼ S. cantoniensis 3 S. trilobata L.)34b. Leaf blades 6 pale green abaxially; fruits usually

well developing (natural species usually not cultivat-ed except in experimental plantations).35a. Leaf blades on sterile long shoots suborbicular,

usually with 1 or 2 pairs of lateral lobes, baseoften rounded or subcordate . . . . . . . S. trilobata L.

35b. Leaf blades on sterile long shoots rhombic-ovate or ovate, often with 2 or 3 pairs of laterallobes, base broadly cuneate . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . S. blumei G. Don (cf. Note 3)

30b. Leaf blades pubescent, sericeous, or tomentose throughout abaxialsurface.36a. Pedicels and hypanthia externally usually glabrous.

37a. Leaf blades on sterile long shoots narrowly elliptic tooblong-lanceolate, pubescent only abaxially . . . . . . . . . . . . .. . . . . . . . . S. cantoniensis var. jiangxiensis (Z. X. Yu) L. T. Lu

37b. Leaf blades on sterile long shoots rhombic-ovate to elliptic,pubescent on both surfaces . . . . . . . . . . . . . . S. pubescens Turcz.

36b. Pedicels and hypanthia externally pubescent to tomentose.38a. Leaf blades pubescent or sericeous abaxially.

39a. Leaf blades light green on both surfaces, mostly 1–2cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nishimurae Kitag.

39b. Leaf blades deep green adaxially and glaucousabaxially, mostly 2–5 cm.40a. Shoots pubescent initially; leaf blades sparsely

pubescent abaxially initially, petioles 5–10 mm;inflorescences . 20-flowered . . . . . . . . . . . . . . .. . . S. hirsuta (Hemsl.) C. K. Schneid. (cf. Note 4)

40b. Shoots densely tomentose initially; leaf bladeslaxly sericeous abaxially at all times, petioles2–3 mm; inflorescences , 20-flowered . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tatakaensis

38b. Leaf blades tomentose abaxially.41a. Shoots initially and leaf blades abaxially yellowish or

rusty yellow tomentose; sepals ovate-lanceolate . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chinensis Maxim.

41b. Shoots initially and leaf blades abaxially grayish towhitish tomentose; sepals triangular to ovate-triangu-lar.42a. Leaf blades on sterile long shoots rhombic-

ovate, serrate, doubly serrate to 6 acutelylobed, with apex usually acute . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . S. dasyantha Bunge (cf. Note 4)

42b. Leaf blades on sterile long shoots broadlyovate, suborbicular, or obovate, shallowlycrenate to incised obtusely lobed and second-arily serrate, with apex obtuse or rounded . . .. . . . . . . . . . . . . . . . . . . . . . . . S. yunnanensis Franch.

TAXONOMIC NOTES FOR CHINESE SPIRAEA inner’’ (Ohwi, 1965: 519; Ikeda, 2001: 104). Thischaracter is reliable and stabilized to variable extent

Note 1. The character of exterior bud scales in in Spiraea species. For instance, the state of twoSpiraea, i.e., two valvate scales versus several valvate bud scales is the conspicuous, stableimbricate ones, has been noted as diagnostic in the character in S. mollifolia, but is not stable in S.literature (e.g., Rehder, 1913, 1927, 1940; Pojarko- flexuosa or S. lichiangensis. Reports for bud scalesva, 1939; Yu & Kuan, 1963), occasionally interpret- are quite confused for S. miyabei Koidz. (species withed as ‘‘outer scales longer versus shorter than the inflorescences compound), where the three known

Volume 21, Number 3 Businsky 3152011 Review of Chinese Spiraea (Rosaceae)

Chinese varieties (var. glabrata Rehder, var. tenui- Note 4. Spiraea nervosa Franch. & Sav. (Enum.folia Rehder, var. pilosula Rehder) are reported to Pl. Jap. 2: 331. 1878) was incorrectly given in Chinesehave small buds covered by several scales, whereas floras (Yu & Lu, 1974; Lu & Crinan, 2003) as abuds of the Japanese variety miyabei were observed synonym to the dissimilar S. dasyantha Bunge (Enum.with two distinctly long, acuminate, valvate scales Pl. Chin. Bor. 23. 1833). The former taxon is accepted(Businsky & Businska, 2002). Generally speaking, as a Japanese endemic (Ikeda, 2001); however, itsthe condition of bud scales is fully developed and the differences from the Chinese S. hirsuta (Bull. Herb.most distinct in mature winter buds, which are Boissier, ser. 2, 3: 342. 1905), examined on limitedusually not present in herbarium specimens nor material and also from published characters, areobserved on plants during vegetation seasons. On the insufficiently convincing, and these taxa could becontrary, buds developing in the leaf axillae on conspecific. In this instance, the name of the Japaneseimmature shoots could be covered initially with taxon would have nomenclatural priority, but a morevalvate scales, which are ripped up later by the thorough comparative study is necessary.increasing bud with several imbricate scales. Acknowledgments. I am grateful to the manage-

Note 2. Spiraea mollifolia was studied by the ment of the Silva Tarouca Research Institute for

present author in several populations from western Landscape and Ornamental Gardening (RILOG),

Hubei (Dashennongjia massif) and southern Shaanxi Pruhonice, Czech Republic, for supporting my

(Qin Ling Mtns.) through western Sichuan and research expeditions to Southeast Asia. The research

northwestern Yunnan to south-central Xizang (Lhasa was also supported by the Ministry of the Environ-

environs). This species is easily recognized by its ment of the Czech Republic (Research Plan grant No.

distinctive buds that are conspicuously long and MZP0002707301). I am indebted to Vıt Vojta and his´narrow, as well as by the thick, angled shoots. wife, Feng-Yun Vojtova, Prague, for their help with

However, it is also extremely variable in its the identification and translation of Chinese texts on

indumentum, which varies from glabrous morphotypes herbarium labels, on maps, and in the literature.

to very densely villous ones, similarly and even more Similar appreciation is given to Wu Zheng-Yi,Institute of Botany, Academia Sinica, Kunming, andso than its sister S. lasiocarpa. For this reason, theto Wu Peng-Cheng, Institute of Botany, Academiaindumentum characters used to discriminate theseSinica, Xiang Shan, Beijing, for their help with thetwo species (the latter under the name S. mongolica)transliteration of the Chinese labels of relevantin the taxonomic key for the Flora of China treatmentherbarium specimens deposited in the herbaria(Lu & Crinan, 2003) is not accepted here.KUN and PE, respectively, and for providing help

Note 3. Spiraea alaschanica Y.-Z. Zhao & T.-J. with field research. I wish especially to thank myWang (Bull. Bot. Res., Harbin 20[4]: 362. 2000) was wife, Ludmila Businska, for many years of her infinitedescribed only from the type, Ningxia Agric. College help, above all in the field during the relevant60009 (holotype, NXAC), reported from Helan Shan research expeditions.(Ningxia, China), 23 May 1960. Because the originalmaterial is limited and not easily accessible, the Literature Citedidentity of this taxon remains unresolved to date. In

Bean, W. J. 1981. Trees and Shrubs Hardy in the Britishthe Flora of China treatment (Lu & Crinan, 2003), it Isles, Vol. 4, 8th ed. [1980, corrected reprint.] Johnwas mentioned as a problematic taxon only in the Murray, London.remark to S. blumei with which it was putatively Businsky, R. & L. Businska. 2002. The genus Spiraea in

related. According to the protologue (Wang & Zhao, cultivation in Bohemia, Moravia and Slovakia. ActaPruhon. 72: 1–165.2000: 362), S. alaschanica corresponds with S.

Chen, I. S. 1978. A new species of the genus Spiraea inblumei except in the reported morphology of buds Taiwan. Formosan Sci. 32(3): 53–55.of the former ‘‘gemmae subulatae, biperulatae.’’ Fu, L.-K. & T. Hong (editors). 2003. Higher Plants ofHowever, this character was assessed by the China, Vol. 6. Qingdao Publishing House, Qingdao.

Grubov, V. I. 1982. Opredelitel’ sosudistych rastenijdescribing authors from young, undeveloped budsMongolii [Key to Vascular Plants of Mongolia]. Nauka,

on material collected at springtime, although the Leningrad.obviously young bud on the accompanying illustra- Handel-Mazzetti, H. 1933. Spiraea L. Pp. 450–454 intion was described as a winter bud. For the reason Symbolae Sinicae, Vol. 7 (Anthophyta), Part 3. J.

that this character is not reliable in some species and Springer, Vienna.Ikeda, H. 2001. Spiraea L. Pp. 104–111 in K. Iwatsuki, D.is fully distinct in mature winter buds (cf. Note 1), S. E. Boufford & H. Ohba (editors), Flora of Japan, Vol. 2b

alaschanica is provisionally accepted here as a (Angiospermae: Dicotyledoneae: Archichlamydeae). Ko-synonym to S. blumei. dansha Ltd., Tokyo.

316 Novon

Institutum Botanicum Kunmingense Academiae Sinicae in tribe Spiraeeae (Rosaceae) inferred from nucleotide(editors). 1984. Index Florae Yunnanensis, Vol. 1. sequence data. Pl. Syst. Evol. 266: 105–118.People’s Publishing House, Kunming. Purohit, K. M. & G. Panigrahi. 1991. The Family Rosaceae

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